Many experimental studies have shown that both the average levels (Hooper et al., 2005; Cardinale et al., 2012; Duffy et al., 2017) and temporal stability (Isbell et al., 2015) of ecosystem functions increase with biodiversity. While the generality of positive relationships between biodiversity and ecosystem functioning is well established (Cardinale et al., 2012; Duffy et al., 2017; Isbell et al., 2015; Lefcheck et al., 2015), current research focuses on the underlying mechanisms (Eisenhauer et al., 2016; Zuppinger-Dingley et al., 2014; Laforest-Lapointe et al., 2017) and context-dependencies (Craven et al., 2016; Guerrero-Ramírez et al., 2017) of biodiversity-ecosystem function relationships, given that ecosystems face progressive environmental changes (Barros and IPCC. Climate Change, 2014). One of the most prominent explanations of why diverse communities perform better than simple ones is that different species complement or facilitate each other in chemical, spatial, and temporal resource use (Loreau and Hector, 2001; Eisenhauer, 2012), resulting in higher functional diversity. Alternatively, diverse communities may have higher functioning because of an elevated probability of containing and becoming dominated by one or a few productive species (Loreau and Hector, 2001). Moreover, there is growing evidence that biotic interactions across trophic levels are important drivers of biodiversity effects (Maron et al., 2011; Schnitzer et al., 2011; Eisenhauer et al., 2012a) and that these interactions are significantly modulated by environmental conditions (Guerrero-Ramírez and Eisenhauer, 2017). As a consequence, biodiversity effects may depend on the environmental context, such as climatic conditions and resource availability (Craven et al., 2016; Guerrero-Ramírez et al., 2017), and thus may change in future environments. However, studies testing biodiversity-ecosystem function relationships under future conditions are rare (Reich et al., 2001; He et al., 2002; Hooper et al., 2012; Thakur et al., 2015), and typically investigate only one or a few ecosystem functions.

Soils provide many ecosystem functions (Bardgett and van der Putten, 2014) that contribute to human well-being, including resources that support plant biomass production, such as nutrients and water, soil carbon storage, and soil erosion control (Wall et al., 2015; Amundson et al., 2015). Consequently, concern about factors that influence soil functioning have attracted increasing scientific (Bardgett and van der Putten, 2014; Amundson et al., 2015; Veresoglou et al., 2015) and public attention (Wall et al., 2015; World Health Organization and Secretariat of the Convention on Biological Diversity, 2014). Changes in plant diversity have been identified as a key factor influencing soil organisms (Hooper et al., 2000; Scherber et al., 2010; Eisenhauer et al., 2013; Lange et al., 2015). Through bottom-up effects, diverse plant communities provide a higher quantity and quality of plant-derived inputs to soil microorganisms and detritivores (Hooper et al., 2000; Eisenhauer et al., 2013), which has cascading effects on the abundance and diversity at higher trophic levels in the soil (Hooper et al., 2000; Scherber et al., 2010; Eisenhauer et al., 2013). Ecosystem functions mediated by soil organisms, such as soil carbon storage (Lange et al., 2015; Fornara and Tilman, 2008), litter decomposition (Vogel et al., 2013), and soil aggregate stabilization (Gould et al., 2016), are thus sensitive to changes in the diversity of plant communities. Given the ecological and economic importance of soil functions (Bardgett and van der Putten, 2014; Wall et al., 2015; Amundson et al., 2015), a key research priority is to understand whether maintaining high local plant diversity secures multiple soil-mediated functions in future environmental conditions.

Ecosystem functioning is inherently multidimensional, and so multifunctionality measures have been increasingly used to summarize the ability of an ecosystem to deliver multiple functions or services simultaneously (Manning et al., 2018). Future environmental conditions may alter the relationship between biodiversity and ecosystem multifunctionality (Manning et al., 2018; Hector and Bagchi, 2007; Soliveres et al., 2016; Byrnes et al., 2014), potentially in distinct ways depending on the response of plant species abundances, or evenness, to new conditions. There is considerable evidence that different plant species promote different ecosystem functions (Hector and Bagchi, 2007), and that different plant species provide different ecosystem functions under different global change scenarios (Isbell et al., 2011). Resource additions often favor ‘fast’ (i.e. acquisitive) plant strategies, thereby decreasing the evenness, functional diversity, and species richness of plant communities (Reich, 2014a). In some cases, the contribution of those fast species to multiple ecosystem functions might not be strong enough to compensate for losses in contribution of sub-dominant species (Allan et al., 2015; van der Plas et al., 2016), thus leading to a reduction in individual functions and multifunctionality. In contrast, if functionally dissimilar plant species complement each other in resource uptake and storage strategies, increased resource availability might not reduce species evenness, and instead could increase productivity and support increased and even provisioning of individual functions (Reich et al., 2001) and multifunctionality. For example, elevated CO₂ concentrations and N addition have been reported to enhance plant diversity effects on plant biomass production (Reich et al., 2001) and soil microbial biomass (Eisenhauer et al., 2013). Due to these contrasting scenarios and given the importance of soils for ecosystem functioning (Bardgett and van der Putten, 2014), evaluating effects of environmental drivers on the multifunctionality of soils and biodiversity-multifunctionality relationships is critically important.

Here, we report root biomass, soil respiration, soil microbial biomass, and water-stable soil aggregates (representing key functions being altered in disturbed soils) (Amundson et al., 2015), correlations among these functions, as well as ecosystem multifunctionality (Hector and Bagchi, 2007; Byrnes et al., 2014) in a long-term grassland plant diversity experiment, with orthogonal manipulation of atmospheric CO₂ concentrations and soil N availability in Minnesota, USA (the BioCON experiment) (Reich et al., 2001). We measured soil functions in 315 plots, 17 years after establishment of the experimental treatments. To calculate ecosystem multifunctionality, we standardized all functions to values ranging between 0 and 1, and then calculated the average level of ecosystem multifunctionality per plot using the mean of the four standardized functions. To evaluate whether average multifunctionality was a result of plant communities simultaneously performing multiple functions at high levels, we determined ecosystem multifunctionality based on the multiple threshold approach, and we more specifically examined four focal performance thresholds (20, 40, 60, and 80%) (Byrnes et al., 2014). The study design enables us to test plant diversity–ecosystem function relationships for multiple soil functions under ambient environmental conditions and under three different global change manipulations that are related to altered resource availability. In addition, we determined treatment effects on realized plant species richness, plant community evenness, the functional diversity of the plant community, and the evenness of multiple soil functions, to explore potential environmental change-induced shifts in the dominance structure, functional diversity, as well as in the even supply of multiple functions (see detailed Materials and methods description below). We find that plant diversity has neutral to positive effects on single soil ecosystem functions and enhances ecosystem multifunctionality. Plant diversity effects on ecosystem multifunctionality are mediated by higher plant community functional diversity and more even provision of multiple soil functions. Although N addition may weaken plant diversity effects on ecosystem multifunctionality at higher thresholds, positive biodiversity-ecosystem functioning relationships were, nevertheless, significant across all the tested global change environments, indicating that conserving local plant diversity is a robust strategy to maintain valuable soil ecosystem services in future environmental conditions.

While it is well established that increased plant diversity (Hector et al., 1999) and resource availability (Lee et al., 2010) enhance the functioning of grasslands, potential interactive effects of these factors are less well studied and are mostly based on responses of single functions like primary productivity (Isbell et al., 2015; Craven et al., 2016; Reich et al., 2001; Stocker et al., 1999). Our study shows the first empirical evidence that significant plant diversity effects on multiple soil functions, and therefore ecosystem multifunctionality, are largely robust to changes in environmental conditions caused by elevated atmospheric CO₂ concentrations, N inputs, and both factors in combination. That is, although there was turnover in the identity of best performing monocultures for each response variable and each global change scenario, the biodiversity effects held in three broad contexts that may occur in the future (Craven et al., 2016; Loreau and Hector, 2001). However, plant diversity effects on ecosystem multifunctionality at higher thresholds of functionality may be impaired under future environmental conditions (Figure 4), potentially by reducing the taxonomic and functional diversity of the plant community (Figure 5). Plant diversity effects on ecosystem multifunctionality were mediated by higher levels of plant community functional diversity and the more even provisioning of multiple soil functions. Our measurements of root biomass, soil respiration, soil microbial biomass C, and percentage of water-stable soil aggregates are indicators of belowground plant biomass production, organic matter decomposition, soil carbon storage (Lange et al., 2015), and soil erosion control, respectively, which denote some of the most crucial ecosystem services grasslands provide for human well-being (Millennium Ecosystem Assessment, 2005). Thus, management policies targeted to maximize these services through maintaining or increasing plant diversity should be applicable under a wide range of contexts (Manning et al., 2018), although high availability of certain resources may compromise beneficial plant diversity effects on high levels of functioning.

Previous studies, including from this field experiment (Reich, 2009), have shown that the composition, species richness, and functioning (e.g. plant biomass production) of grasslands can be significantly affected by anthropogenic environmental changes, such as climate change (Reich et al., 2014b; Grime et al., 2000), elevated atmospheric CO₂ concentrations (Reich et al., 2001; Morgan et al., 2011), and N deposition (Clark and Tilman, 2008; Simkin et al., 2016). In contrast, in the present study, elevated CO₂ concentrations and N input had no significant effects on soil functions (Table 1), despite significantly increased aboveground plant biomass in the N addition treatment (Table 2). These overall weak or missing effects might be because the response of grassland functions to environmental change depends on local conditions like precedent climate events and history of anthropogenic disturbances (Grime et al., 2000; Simkin et al., 2016), although we did not record any particularly harsh environmental conditions in the year of the study. Previous research in the BioCON experiment showed significant CO₂ and N effects on plant biomass (Reich et al., 2001) and total belowground carbon allocation, as well as on soil communities (He et al., 2010; Eisenhauer et al., 2012b) and functions (Chung et al., 2007; Reich et al., 2018). However, the effect of those global change drivers on total belowground carbon allocation and soil functions may increase or decrease over time and vary among years and species compositions alone or jointly (Reich et al., 2018; Adair et al., 2009; Andresen et al., 2016). The long-term character of the BioCON experiment (17 y of experimental treatments in the sampling year) (Reich et al., 2001), the gradual increase of biodiversity effects over time (Reich et al., 2012), and the fact that some soil functions, such as root biomass and soil aggregate stability, should reflect treatment effects of multiple years, suggest that the present results are unlikely to be due to the fact that they are based on a single sampling campaign. This is supported by previous assessments of soil respiration (Eisenhauer et al., 2013), soil microbial biomass (Eisenhauer et al., 2013), and root biomass (Reich et al., 2001) that showed significant positive plant diversity effects. However, environmental change effects may be more important if higher thresholds of functionality are considered (Figure 4), showing that the multiple thresholds approach can provide additional insights into the functioning of ecosystems.

In contrast to the effects of the other two global change drivers, our study showed that plant diversity had strong and consistent effects on three out of the four single soil functions we investigated, as well as on ecosystem multifunctionality. In line with other studies (Reich et al., 2012; Mueller et al., 2013; Ravenek et al., 2014), we observed positive effects of plant diversity on root biomass. High-diversity plant communities are more likely to include different root traits, that is deep- and shallow-rooting species, than low-diversity communities (Mueller et al., 2013). This may lead to a more complete usage of the available habitat space, thus increasing total resource extraction from the soil (Ravenek et al., 2014; Mommer et al., 2010). Thereby, higher diversity plant communities have been shown to increase root biomass in surface soils (Ravenek et al., 2014) and the proportion of deep roots (Mueller et al., 2013). Increased plant biomass production at high plant diversity (above and below the ground) and increased diversity of organic inputs to the soil have been shown to enhance soil microbial biomass and activity (Hooper et al., 2000; Eisenhauer et al., 2013; Lange et al., 2015). This was also found in the present study (Tables 1 and 2, Figure 1b,c) and indicates higher decomposition activity and turnover of microbial products with potential consequences for soil carbon storage (Lange et al., 2015). The simultaneous increase in functions related to soil carbon storage (e.g. soil microbial biomass and root biomass) and decomposition activity (soil respiration) may appear counterintuitive as one may expect trade-offs between these functions. However, these results are in line with recent findings showing that elevated microbial activity, such as indicated by soil respiration, reflects the transfer of higher amounts of plant products into microbial products that then accumulate in the soil (Lange et al., 2015; Cotrufo et al., 2015). Thus, although high soil respiration means greater short-term losses of organic matter as CO₂, it can also indicate enhanced long-term soil carbon storage (Lange et al., 2015; Cotrufo et al., 2015) as microbial necromass might end up in slow-cycling soil organic matter pools (Cotrufo et al., 2015; Schmidt et al., 2011).

In contrast to previous studies, the proportion of water-stable soil aggregates did not significantly increase with plant diversity (Gould et al., 2016; Pérès et al., 2013), root biomass (Gould et al., 2016), microbial biomass (Gould et al., 2016; Pérès et al., 2013), elevated CO₂ (Rillig et al., 2001), or N addition (Riggs et al., 2015), although positive trends were observed with diversity (Figure 1d). Soil aggregate stability contributes to the regulating ecosystem service soil erosion control (Millennium Ecosystem Assessment, 2005), which is of major relevance in a rapidly changing world with increased likelihood of extreme weather events (Amundson et al., 2015; Fischer et al., 2018). Soil aggregate stability also informs soil carbon storage, as aggregates incorporate organic matter protecting it from decomposition (Six et al., 2000) on decadal timescales. Although soil aggregate stability depends on soil biological properties, such as (fine) root biomass and root length density (Gould et al., 2016), and soil biota including microbes and animals (Lehmann et al., 2017), abiotic factors like soil texture are also important controls (Denef et al., 2002) and may have prevailed in the very sandy soil of the present study (Reich et al., 2001).

Our study also showed the importance of plant diversity for the simultaneous provisioning of multiple functions (soil multifunctionality). Despite the increasing awareness of the importance of biodiversity for ecosystem multifunctionality (Lefcheck et al., 2015; Manning et al., 2018; Hector and Bagchi, 2007; Soliveres et al., 2016; Byrnes et al., 2014), knowledge of how these results hold under future environmental conditions is limited. Increases in N deposition and in CO₂ levels are predicted with future global change and represent elevated resource availability. Elevated resources can facilitate the dominance of certain species that are more efficient in resource use leading to reduced taxonomic and functional diversity of plant communities (Morgan et al., 2011; Clark and Tilman, 2008; Simkin et al., 2016). This, in turn, could be expected to offset the observed positive effects of plant diversity on multiple ecosystem functions (Hector and Bagchi, 2007; Soliveres et al., 2016; Byrnes et al., 2014). Indeed, elevated resource availability decreased realized plant species richness, community evenness, and functional diversity in the present study (Table 2, Figure 4—figure supplements 1–5). However, our results indicate that higher sown plant diversity attenuated the detrimental effects of elevated resource availability on realized plant diversity indices, and we found that positive effects of plant diversity on single ecosystem functions, as well as on soil multifunctionality, were largely independent of environmental conditions. Although these results were robust to different analytical approaches to quantify multifunctionality (Byrnes et al., 2014), and also hold in realistic scenarios where plant diversity loss is relatively moderate (Eisenhauer et al., 2013) (excluding monocultures), plant diversity effects on ecosystem multifunctionality were attenuated for high thresholds (Byrnes et al., 2014) under changed environmental conditions. Thus, our study provides strong evidence that the positive effects of plant diversity on soil functioning are not only applicable at present, but also in realistic future global change scenarios. However, our results also indicate that environmental conditions decreasing the functional diversity of plant communities may threaten the multifunctionality of soils.

The present and other grassland studies have shown that biodiversity effects on aboveground functions (aboveground productivity) can also be robust to changes in N addition and water availability (Craven et al., 2016). This contrasts with other findings on context-dependent biodiversity effects (Guerrero-Ramírez et al., 2017; Ratcliffe et al., 2017). Future studies should investigate the causes of these inconsistent findings, for example by considering variations in the experimental age (Craven et al., 2016; Guerrero-Ramírez et al., 2017; Thakur et al., 2015), local biotic and abiotic conditions (Guerrero-Ramírez et al., 2017; Ratcliffe et al., 2017), and the focal functions (Byrnes et al., 2014; Allan et al., 2015). While the effects of biodiversity on certain ecosystem functions in certain habitat types may become stronger or weaker under future conditions, we show that grassland biodiversity effects on at least the belowground functions we studied are robust to environmental change as long as high levels of plant functional diversity are maintained. Thus, studying shifts in the functional composition of plant communities in response to environmental change is a promising approach to better understand the context dependency of biodiversity effects based on species richness.

In summary, our study showed strong and consistent effects of plant diversity on various belowground ecosystem functions related to biomass production, nutrient cycling, and carbon sequestration, as well as on ecosystem multifunctionality. These effects were much stronger than those of two other global change drivers, stressing the pivotal importance of plant diversity (Hooper et al., 2012; Thakur et al., 2015) for soil functioning. The present study sheds light on the underlying mechanisms of plant diversity effects on ecosystem multifunctionality by stressing the role of plant community functional diversity as well as the more even provisioning of multiple ecosystem functions at high plant diversity. Importantly, we also show that these positive effects of biodiversity were largely robust to environmental changes in N deposition and CO₂ levels, indicating that biodiversity will be equally important under future conditions as at present. Notably, high levels of soil multifunctionality may be threatened by environmental change-induced reductions in the functional diversity of plant communities.

All data generated or analyzed during this study are available in Supplementary File 3.

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Author details

1. Nico Eisenhauer

   1. German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany
   2. Institute of Biology, Leipzig University, Leipzig, Germany

   Contribution

   Conceptualization, Resources, Data curation, Formal analysis, Supervision, Funding acquisition, Investigation, Visualization, Writing—original draft, Project administration

   Contributed equally with

   Jes Hines

   For correspondence

   nico.eisenhauer@idiv.de

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-0371-6720

2. Jes Hines

   1. German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany
   2. Institute of Biology, Leipzig University, Leipzig, Germany

   Contribution

   Conceptualization, Formal analysis, Investigation, Visualization, Writing—original draft

   Contributed equally with

   Nico Eisenhauer

   Competing interests

   No competing interests declared

3. Forest Isbell

   Department of Ecology, Evolution, and Behavior, University of Minnesota, St Paul, United States

   Contribution

   Writing—review and editing

   Competing interests

   No competing interests declared

4. Fons van der Plas

   Department of Systematic Botany and Functional Biodiversity, Leipzig University, Leipzig, Germany

   Contribution

   Writing—review and editing

   Competing interests

   No competing interests declared

5. Sarah E Hobbie

   Department of Ecology, Evolution, and Behavior, University of Minnesota, St Paul, United States

   Contribution

   Writing—review and editing

   Competing interests

   No competing interests declared

6. Clare E Kazanski

   Department of Ecology, Evolution, and Behavior, University of Minnesota, St Paul, United States

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-7432-5666

7. Anika Lehmann

   1. Institute of Biology, Freie Universität Berlin, Berlin, Germany
   2. Berlin-Brandenburg Institute of Advanced Biodiversity Research, Berlin, Germany

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

8. Mengyun Liu

   1. German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany
   2. Institute of Biology, Leipzig University, Leipzig, Germany
   3. Key Laboratory of Vegetation and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China
   4. University of Chinese Academy of Sciences, Beijing, China

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

9. Alfred Lochner

   1. German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany
   2. Institute of Biology, Leipzig University, Leipzig, Germany

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

10. Matthias C Rillig

    1. Institute of Biology, Freie Universität Berlin, Berlin, Germany
    2. Berlin-Brandenburg Institute of Advanced Biodiversity Research, Berlin, Germany

    Contribution

    Writing—review and editing

    Competing interests

    No competing interests declared

11. Anja Vogel

    1. German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany
    2. Institute of Biology, Leipzig University, Leipzig, Germany
    3. Institute of Ecology and Evolution, Friedrich Schiller University Jena, Jena, Germany

    Contribution

    Data curation, Formal analysis, Writing—review and editing

    Competing interests

    No competing interests declared

12. Kally Worm

    Department of Forest Resources, University of Minnesota, St Paul, United States

    Contribution

    Investigation, Writing—review and editing

    Competing interests

    No competing interests declared

13. Peter B Reich

    1. Department of Forest Resources, University of Minnesota, St Paul, United States
    2. Hawkesbury Institute for the Environment, Western Sydney University, Sydney, Australia

    Contribution

    Resources, Supervision, Investigation, Writing—review and editing

    Competing interests

    No competing interests declared

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